Stephen R. Ross

Hand preferences for coordinated bimanual actions in 777 great apes: Implications for the evolution of handedness in Hominins

Whether or not nonhuman primates exhibit population-level handedness remains a topic of considerable scientific debate. Here, we examined handedness for coordinated bimanual actions in a sample of 777 great apes including chimpanzees, bonobos, gorillas, and orangutans. We found population-level right-handedness in chimpanzees, bonobos and gorillas, but left-handedness in orangutans. Directional biases in handedness were consistent across independent samples of apes within each genus. We suggest that, contrary to previous claims, population-level handedness is evident in great apes but differs among species as a result of ecological adaptations associated with posture and locomotion. We further suggest that historical views of nonhuman primate handedness have been too anthropocentric, and we advocate for a larger evolutionary framework for the consideration of handedness and other aspects of hemispheric specialization among primates.

Comparative Assessment of Handedness for a Coordinated Bimanual Task in Chimpanzees (Pan troglodytes), Gorillas (Gorilla gorilla), and Orangutans (Pongo pygmaeus)

Hand preferences for a coordinated bimanual task were assessed in a sample of 31 captive gorillas (Gorilla gorilla) and 19 captive orangutans (Pongo pygmaeus) and were compared with chimpanzee (Pan troglodytes) hand preferences in subjects that were matched on the basis of age, sex, and rearing history. The task required that the apes remove food from the inside edges of a symmetrical polyvinyl chloride pipe presented to them in their home cages. The results indicate significant species differences with chimpanzees showing population-level right-handedness and orangutans showing population-level left-handedness. The gorillas showed a nonsignificant trend toward right-handedness. The results are discussed in terms of possible ecological or biomechanical factors that may influence hand preferences in different ape species.

Primate energy expenditure and life history

Significance Measurements of daily energy expenditure indicate that primates, including humans, expend only half of the calories expected for mammals of similar body size. As energy expenditure is central to organismal biology, these results hold important implications for life history, evolutionary biology, and foraging ecology for primates and other mammals. Specifically, we show that primates’ remarkably low metabolic rates account for their distinctively slow rates of growth, reproduction, and aging. Humans and other primates are distinct among placental mammals in having exceptionally slow rates of growth, reproduction, and aging. Primates’ slow life history schedules are generally thought to reflect an evolved strategy of allocating energy away from growth and reproduction and toward somatic investment, particularly to the development and maintenance of large brains. Here we examine an alternative explanation: that primates’ slow life histories reflect low total energy expenditure (TEE) (kilocalories per day) relative to other placental mammals. We compared doubly labeled water measurements of TEE among 17 primate species with similar measures for other placental mammals. We found that primates use remarkably little energy each day, expending on average only 50% of the energy expected for a placental mammal of similar mass. Such large differences in TEE are not easily explained by differences in physical activity, and instead appear to reflect systemic metabolic adaptation for low energy expenditures in primates. Indeed, comparisons of wild and captive primate populations indicate similar levels of energy expenditure. Broad interspecific comparisons of growth, reproduction, and maximum life span indicate that primates’ slow metabolic rates contribute to their characteristically slow life histories.

Metabolic acceleration and the evolution of human brain size and life history

Humans are distinguished from the other living apes in having larger brains and an unusual life history that combines high reproductive output with slow childhood growth and exceptional longevity. This suite of derived traits suggests major changes in energy expenditure and allocation in the human lineage, but direct measures of human and ape metabolism are needed to compare evolved energy strategies among hominoids. Here we used doubly labelled water measurements of total energy expenditure (TEE; kcal day−1) in humans, chimpanzees, bonobos, gorillas and orangutans to test the hypothesis that the human lineage has experienced an acceleration in metabolic rate, providing energy for larger brains and faster reproduction without sacrificing maintenance and longevity. In multivariate regressions including body size and physical activity, human TEE exceeded that of chimpanzees and bonobos, gorillas and orangutans by approximately 400, 635 and 820u2009kcal day−1, respectively, readily accommodating the cost of humans’ greater brain size and reproductive output. Much of the increase in TEE is attributable to humans’ greater basal metabolic rate (kcal day−1), indicating increased organ metabolic activity. Humans also had the greatest body fat percentage. An increased metabolic rate, along with changes in energy allocation, was crucial in the evolution of human brain size and life history.

Brief communication: Captive gorillas are right-handed for bimanual feeding.

Predominance of right-handedness has historically been considered as a hallmark of human evolution. Whether nonhuman primates exhibit population-level manual bias remains a controversial topic. Here, we investigated the hypothesis that bimanual coordinated activities may be a key-behavior in our ancestors for the emergence and evolution of human population-level right-handedness. To this end, we collected data on hand preferences in 35 captive gorillas (Gorilla gorilla) during simple unimanual reaching and for bimanual coordinated feeding. Unimanual reaching consisted of grasping food on the ground, while bimanual feeding consisted of using one hand for holding a food and processing the food item by the opposite hand. No population-level manual bias was found for unimanual actions but, in contrast, gorillas exhibited a significant population-level right-handedness for the bimanual actions. Moreover, the degree of right-handedness for bimanual feeding exceeds any other known reports of hand use in primates, suggesting that lateralization for bimanual feeding is robust in captive gorillas. The collective evidence is discussed in the context of potential continuity of handedness between human and nonhuman primates.

An experimental, comparative investigation of tool use in chimpanzees and gorillas

Studies of ape tool use have been conducted in captivity since the early 1900s and in the wild since the 1960s. Chimpanzees are the most prolific tool users among the apes, and are known to use more tools than any other nonhuman animal. In contrast, reports of gorilla tool use are rare both in wild and captive settings. Studies of the processes involved in tool use learning have been limited in the wild by the lack of ability to control several unpredictable variables, and in captivity by tool use opportunities that are often presented in non-naturalistic contexts. We attempted to address both of these limitations by providing naive subjects with a naturalistic tool use device (built to simulate a termite mound) while housed in a more natural social setting to approximate how learning would occur in the wild. Both gorillas and chimpanzees participated in the experiment to allow comparative analyses of acquisition of tool behaviour and the factors that may affect acquisition. Both species showed low frequencies of interaction with the mound in the baseline condition, before baiting with a food reward. Once baited, chimpanzees both attempted and succeeded to extract the reward more quickly than did gorillas. The number of social group members at the mound was significantly higher for chimpanzees than for gorillas and may have affected skill acquisition. We advocate that comparative approaches to skill acquisition and learning are valuable, but that researchers need to be cognizant of species differences in social structure that may affect results.

Issues of choice and control in the behaviour of a pair of captive polar bears (Ursus maritimus).

Stereotyped behaviour occurs in a wide variety of captive animals including ursids. The provision of animal control over aspects of their environment by providing choices is a critical element for improving welfare. The behaviour of two sibling polar bears at a metropolitan zoo was examined to investigate the effect of providing access to their indoor, off-exhibit holding space. Both bears demonstrated behavioural changes when given the choice to access their indoor dens including decreased stereotyped behaviours and increased social play. These results, although based on just two bears, provide additional support for the assertion that choice and control are closely tied to issues of well-being for captive animals.

Influences on visitor behavior at a modern immersive zoo exhibit.

Zoos serve as centers for both research and education. The challenge is to convey messages about their conservation projects while meeting visitor expectations, which often include recreation and entertainment. One way this can be achieved is through the design of immersive exhibits that draw visitors in and engage them with interactive educational elements. Regenstein African Journey (RAJ) opened at Lincoln Park Zoo in 2003 and was designed to take visitors on a simulated safari through Africa. Because visitor experience was a major design goal, we conducted a timing and tracking study to evaluate use of the building and educational components. For a 9-week period in 2003, we tracked 338 visitors to RAJ and recorded continuous data as they moved through the building. Data were collected on handheld computers that provided precise timing data. The median visit was 11.08 min, 41% of which was spent looking at animals and 9% of which was spent engaged with interpretive elements. We found significant differences in the way visitors used signage: those in groups without children spent more of their visit engaged with signage than those with children and visitors who spent more of their visit interacting socially spent less time engaged with signage. By understanding how visitors use the educational opportunities presented to them, we can better meet their expectations and more effectively achieve the goal of conservation education.

Inappropriate Use and Portrayal of Chimpanzees

Depictions of chimpanzees as caricatures can lead people to think these animals are not endangered and is a problem for conservation and welfare efforts.

Specific Image Characteristics Influence Attitudes about Chimpanzee Conservation and Use as Pets

Chimpanzees are endangered in their native Africa but in the United States, they are housed not only in zoos and research centers but owned privately as pets and performers. In 2008, survey data revealed that the public is less likely to think that chimpanzees are endangered compared to other great apes, and that this is likely the result of media misportrayals in movies, television and advertisements. Here, we use an experimental survey paradigm with composite images of chimpanzees to determine the effects of specific image characteristics. We found that those viewing a photograph of a chimpanzee with a human standing nearby were 35.5% more likely to consider wild populations to be stable/healthy compared to those seeing the exact same picture without a human. Likewise, the presence of a human in the photograph increases the likelihood that they consider chimpanzees as appealing as a pet. We also found that respondents seeing images in which chimpanzees are shown in typically human settings (such as an office space) were more likely to perceive wild populations as being stable and healthy compared to those seeing chimpanzees in other contexts. These findings shed light on the way that media portrayals of chimpanzees influence public attitudes about this important and endangered species.